The Scuttle Fly (Diptera: Phoridae) Assemblages of Pine Plantations of the Biala Forest (Poland)

The scuttle fly (Diptera: Phoridae) assemblages of pine plantations of the Biala Forest (Poland)

Ewa Durska

Durska, E. 2009: The scuttle fly (Diptera: Phoridae) assemblages ofpine plantations of the Biala Forest (Poland). — Entomol. Fennica 20: 170—178.

Scuttle fly diversity in disturbed habitats was evaluated on plots in pine plantations ofPoland’s Biala Forest. The assemblages present in the two years were as- sessed for the abundance of species, dominance structure, similarity and species richness, as well as in regard to indices ofspecies diversity, evenness and fidelity. Megaselia brevicostalis was the first dominant on each of the three plots and in both study years. The number of species and their abundance was greatest on the turn ofAugust and September. Most ofthe dominants and characteristic species are multivoltine, showing spring and late summer/autumn activity.

E. Darska, Department ofEcology andBiodiversity, Museum andInstitute ono— ology, PAS, Wilcza 64, 00—679 Warsaw, Poland; Email: edarska@miiz. waw.pl Received: 2 April 2008, accepted 5 Jane 2008

1. Introduction strongly to habitat pollution. An increase in pollution brings about significant changes in the pro- Phoridae (scuttle flies) constitute one ofthe most portion of trophic groups and the number of spe- abundant and diverse families of Diptera of cies with zoophagous larvae decreases while that woodlands, meadows, agroecosystems and urban of saprophagous increases (Durska 1996). habitats. To date, over 3200 species have been de- During my studies in the BialowieZa Primeval scribed, a quarter of these from the Palaearctic Forest I also stated that scuttle fly assemblages Region (Disney 1991). might be regarded not only as highly indicatory As recent studies make clear, assemblages of because oftheir sensitivity to habitat changes but scuttle flies not only offer an effective representa- also ofpossessing specific groups of characteris- tion of biodiversity, but also serve in investiga- tic species in all four age-classes (pine plantation, tions of habitat recolonisation (Disney 1994; young-growth stand, timber stand and old- Durska 2001a, 2002, 2003, 2006, Prescher et al. growth stand) (Durska 2001a). 2002; Disney & Durska 2008). It is known that “a complex ecosystem offers In my previous studies, I found that the scuttle more options than a simpler one when placed un- fly assemblage in pine plantations greatly dif- der stress” (Moore 2005), and habitat heterogene- fered in its species composition from the assem- ity due to such disturbances as cutting, grazing, blages in the older age-classes of the succession wildfires or windthrow may create very similar series of moist pine forest in the BialowieZa Pri- conditions for patch-assemblages of insects as meval Forest. I observed a complete exchange of within a patch surrounded by older-phase stands dominant species even in young-growth (Durska (Tschamtke & Brandl 2004). Indeed, disturbance 2001a) may be one of the factors controlling species di- The scuttle fly assemblages also react very versity (Ostman et al. 2006; Tanner & Belling- ENTOMOL. FENNICA Vol. 20 ° Phoridae ofthe Biaia Forest, Poland 171 ham 2006). There is little information on how dis- in 1986. Distances between the plots did not ex- turbances affect the diversity of Phoridae in the ceed 5 km. earliest phase of succession. Only my investiga- Scuttle flies were collected using yellow plas- tions (Durska 2001a, 2006) in the pine planta- tic pans, 18 cm in diameter, containing water so- tions in the BialowieZa Primeval Forest and the lution of75% ethylene glycol for conservation of Tuchola Forest, and those by Prescher et al. the insects and some detergent (Bankowska & (2002) in the chestnut belt of the Alps after Garbarczyk 1982). Flies were thus sampled using wildfires, provides comparable data. five such traps installed (dispersed) on the ground The aim ofmy work was thus to determine the on each plot. Trapping continued from April structure ofthe assemblages of scuttle flies found through to October, with traps emptied fort- on small plots in pine plantations established fol- nightly. lowing clear—cutting in the moist pine forest habi- The use of the yellow plastic pans provides tats ofBiala Forest in Poland, as well as to docu- for the collection of material from both the can- ment the attendant phenology. opy and forest floor (Durska 1996, 2001a). Dis- In this paper, I examine the abundance of spe- ney (2004) reported yellow-trap selectivity for cies, dominance structure, similarity and species particular scuttle fly species while it is known that richness, as well as species diversity, evenness colour (from red via orange to yellow) can manip- and habitat fidelity (considering the level ofsimi- ulate “the direction of flies’ movements and as- larity of assemblages in stands in the old-growth sists the prolongation of the search for food” phase). (Lunau et al. 2005). Overall, studies using yellow pans by myself and others, confirm that ca. 70% of all Phoridae caught are flies of the genus 2. Materials and methods Megaselia (Disney 1994, Durska 1996, 2001a, 2006). In turn, studies by Goos (1975) on flying 2.1. Study area and sampling insects in sugar-beet plantations combine with my own investigations (Durska 1996, 2001a, This study formed a part of a wider research on 2006) to confirm the status ofthe Phoridae among the secondary succession of scuttle fly assem- the families ofDiptera caught most abundantly in blages in moist pine forest habitat, and was car- yellow pans. ried out in Biala Forest in Poland (52°30’—53°00’ N, 20°40’—21°30’ E) in 1986 and 1987. This is a large forest complex (over 64,000 ha) on the P01- 2.2. Statistical analyses ish Lowland with a preponderance of managed tree stands. Plant communities of moist pine fo- I confined my analysis of scuttle fly assemblages rest habitat are represented there by the associa- to the male individuals of all genus present, as tion Peucedano—Pinetum Mat, in its Sarmatian most Megaselia females are not identifiable to variant (Matuszkiewicz et al. 1993). species level at our present state of knowledge According to phytosociological analyses in and so have only been identified to genus level. the area there are slight differences in floristic In describing the assemblages, I took account composition between old-growth and clearings of the relative abundance of males of different with young tree seedlings (Matuszkiewicz et al. species, as well as the dominance structures, per- 1993). Under natural conditions, Scots pine is forming calculations for each plot and each study eliminated by Norway spruce, which shades the year. In terms of the male individuals, “Domi- understorey severely. Clear cutting is therefore nant” species were those with a relative abun- employed commonly to facilitate the regenera- dance 2 1%. tion of pine from seed. I calculated the Shannon-Weaver diversity Three plots (ca. 5 ha) were randomly selected index (H) using the data from each plot, in accor- within even-aged pine plantations. The plots dance with the formula: (coded 62g, 34fand 3271) had either two-year-old pines (34f) or three-year—old pines (62g and 32m) ’=—Zpilogepi (1) 172 Durska ' ENTOMOL. FENNICA Vol. 20 where pi = Ni/N; Ni is the number of individuals F: a/b >< 100 (5) ofthe i-th species and N the total number of individuals. The diversity index (H 3 was calcutated where a = the number of individuals of a given for each study year and the differences between species in a given habitat, and b = the total num- the years were tested using x_-test (Magurran ber of individuals of a given species in all com- 1988) pared habitats. To assess the habitat fidelity of as- My analysis for evenness involved the for- semblages to the study area, I compared the scut- mula: tle fly assemblages of pine plantations (three plots) with those ofBiala Forest stands in the old- J’ = H ’/loges (2) growth phase (also three plots, previous data; Durska 1996), five yellow pans being hung up on where s is the species richness (species number). the crowns of trees on each, during the same two I also applied Spearman rank correlation co- study seasons (1986 and 1987, simultaneous trap- efficients (rs) in comparing the scuttle fly assem- ping period in pine plantations and old-growth blages from the two years. I made further be- stands). I trapped from April through to October, tween-year comparisons of the assemblages us- emptying traps fortnightly at the same time in the ing two similarity indices: Sorensen’s quotient two habitats compared (pine plantations and old- (So) (Sorensen 1948) and Morisita’s quotient growth stands in 1986 and 1987) (Durska 1996). (M0) as modified by Horn (1966). The former is I also compared the seasonal and inter-annual based on presences and absences of species: changes in the abundance (number of individuals) and species richness (number of species) of SQ = 2a/b+c (3) the scuttle fly assemblages from 1986 and 1987. where: a is the number of common species, b is the number of species of the first assemblage 3. Results (1986) and c is the number of species of the sec- ond assemblage (1987). IusedM0 to calculate the I collected a total of 6155 adult individuals of similarity ofdominance structures in respect to all phorid flies (3506 in 1986 and 2649 in 1987), at- species or dominants (abundance 2 1%) in the tributing these to 52 scuttle fly species (42 in 1986 pine plantation assemblages obtained in the two and 44 in 1987), and thus obtaining almost 50% years. The formula is: of the Phoridae known from pine plantations on the Polish Lowland (Disney 1991, Durska 2001a, 2ixiyi b, Durska 2006). (Tables 1—2). The number ofre- i:1 M0 = (4) corded species varied from 23 to 31 in the study ixaz+iyaz plots. 51:1 51:1 where: xi, yl. are the percentages of common species, xa, ya are the percentages of particular spe- 3.1. Assemblage structure cies in each of the assemblages compared (1986 and 1987) ands is the total number of species. M0 In both years, individuals of species in the genus presents the similarity ofdominance structures of Megaselia accounted for over 90% of the phorid high-abundance species very well. This formula assemblage associated with each plot. I confined is appropriate where the data are expressed as the my considerations to males identified to species proportions of common species (xi, y) of the re- level (i.e. 291 1 individuals), these constituting ca. spective samples composed of species 1' (Horn 50% of all specimens collected (Table 1). 1966) I obtained very high values for the indices of I used habitat fidelity (F), expressed quantita- qualitative similarity (SQ) and quantitative simi- tively, to determine those species that were char- larity (M0) calculated for common dominants in acteristic of pine plantations, employing the for- the two seasons (Table 3). mula: The fact that 31 of the 52 species occurred in ENTOMOL. FENNICA V01. 20 ° Phoridae 0fthe Biaia Forest, Poland 173

Table 1. List of scuttle fly (Phoridae) species with numbers of individuals (N) and relative abundances (%I\I) given for the individual years. *: Dominant species with dominance 2 1% during one or both years.

1 986 1 987

Species N %N N %N

Anevrina thoracica (Becker, 1901 —— 1 0.04 Anevrina unispinosa (Zetterstedt, 1860) 1 0.03 —— *Borophaga subsultans (Linnaeus,1767) 10 0.29 —— Conicera florico/a Schmitz, 1938 —— 1 0.04 *Conicera similis (Haliday,1833) 122 3.48 33 1.25 *Diplonevra funebris (Meigen, 1830) 25 0.72 13 0.49 Gymnophora nigripennis Schmitz, 1926 1 0.03 1 0.04 Gymnophora sp. 4 0.11 1 0.04 Megaselia afflnis (Wood, 1909) 2 0.06 —— *Megase/ia altifrons (Wood,1909) 18 0.51 20 0.76 Megaselia basispinata (Lundbeck, 1920) —— 1 0.04 *Megase/ia brevicosta/is (Wood, 1910) 515 14.69 452 17.06 Megaselia campestris (Wood, 1908) 2 0.06 —— Megaselia dah/i (Becker,1901) 3 0.09 1 0.04 *Megase/ia diverse (Wood, 1909) 9 0.26 1 0.04 *Megaselia giraudii—complex (Egger,1862) 21 0.60 48 1.81 Megaselia groen/andica (Lundbeck,1901) 1 0.03 —— Megaselia gregaria (Wood, 19100) —— 1 0.04 Megaselia hemydisneyi Durska, 1998 —— 1 0.04 *Megaselia hya/ipennis (Wood, 1912) 13 0.37 2 0.08 *Megaselia involuta (Wood, 1910) 9 0.26 10 0.38 Megaselia lata (Wood, 1910) —— 1 0.04 Megaselia latifrons (Wood, 1910) 3 0.09 3 0.11 Megaselia longicostalis (Wood, 1912) 2 0.06 2 0.08 Megaselia mal/ochi (Wood, 1909) —— 13 0.49 *Megase/ia manicata (Wood, 1910) 24 0.68 60 2.27 Megaselia meconicera (Speiser, 1925) 1 0.03 1 0.04 *Megaselia minor (Zetterstedt, 1848) 34 0.97 30 1.13 *Megase/ia nigriceps (Loew, 1866) 64 1.83 44 1.66 *Megase/ia pleura/is (Wood, 1909) 61 1.74 33 1.25 Megaselia plurispinulosa (Zetterstedt, 1860) 1 0.03 4 0.15 Megaselia posticata (Strobl, 1898) 1 0.03 —— Megaselia propinqua (Wood, 1909) —— 5 0.15 *Megaselia pulicaria—complex (Fallén, 1823) 123 3.51 112 4.23 *Megaselia pumila (Meigen, 1830) 24 0.68 26 0.98 Megaselia pusil/a (Meigen, 1830) 4 0.11 1 0.04 *Megase/ia scute/Iaris (Wood, 1909) 68 1.94 200 7.55 Megaselia spinigera (Wood, 1908) 1 0.03 —— Megaselia stigmatica (Schmitz, 1920) 1 0.03 —— *Megaselia subnudipennis (Schmitz, 1919) 14 0.04 —— *Megaselia unico/or(Schmitz, 1919) 31 0.88 1 0.04 *Megase/ia verra/Ii (Wood, 1910) 233 6.65 42 1.59 Megaselia woodi (Lundbeck, 1922) 6 0.17 1 0.04 *Megaselia xanthozona (Strobl, 1892) 53 1.51 31 0.79 Megaselia zonata (Zetterstedt, 1838) —— 2 0.08 Metopina heselhausi Schmitz, 1914 1 0.03 5 0.19 *Metopina oligoneura (Mik, 1867) 72 2.05 126 4.76 Phalacrotophora sp. n —— 1 0.04 Triphleba bifida Schmitz, 1949 —— 2 0.08 Triphleba distinguenda (Strobl, 1892) —— 1 0.04 Triphleba minuta (Fabricius, 1787) —— 1 0.04 Triphleba opaca (Meigen, 1830) 1 0.03 1 0.04 Triphleba subcompleta Schmitz, 1927 1 0.03 1 0.04 *Triphleba trinervis (Becker, 1901) 11 0.31 —— Triphleba sp. 1 0.03 —— Megaselia sp. females 1,841 52.51 1,238 46.73 Megaselia sp. males 73 2.08 78 2.94 Megaselia sp. A —— 5 0.19

Total 3,506 100 2,649 100

176 Durska ' ENTOMOL. FENNICA Vol. 20

es, since indentification of many Megaselia rest at Tyresta near Stockholm in August 1999, or males (let alone females) proves problematical. else obtained using ground eclectors located in When comparing the species diversity of the the chestnut belt following a wildfire in the Swiss scuttle fly assemblages from plots in pine planta- Alps, again accounted for over 90% of all tions of three forest areas on the Polish Lowland Phoridae sampled (Prescher et al. 2002; Durska (almost the same age, at the same time and using unpubl.). the same method), I noted the highest diversity Overall, two of the dominants in my study of for the BialowieZa Primeval Forest (1986: H’ = the scuttle fly assemblages on each pine-planta- 3.95, 1987: H’ = 3.86), and the lowest values for tion plot (i.e. the pyrophilous Megasell'a verralli the Tuchola Forest (1986: H’ = 3.05, 1987: H’ = and the polysaprophagous M. brevicostalis) are 2.78). The latter one is a managed forest where also dominants in heterogeneous habitats afflic- the situation in 1987 was affected by the ex- ted by wildfires, as in the aforementioned chest- tremely marked dominance of the polyphagous nut—belt ofthe Swiss Alps and the hemiboreal fo- Megaseliapulicaria — complex, probably as a re- rests ofTyresta (Prescher et al. 2002; Durska un- sult of chemicals being employed against publ.). I also found these same two species to be Neodiprion sertl'fer in 1986 (Durska 2006). The the dominants in the Piska Forest (Poland), in the structure of the scuttle fly assemblages is thus wake of a windthrow incident that took place in seen to be markedly susceptible to quantitative July 2002 (Durska et al. in prep.). environmental differences (Goos 1975; Durska The work I recount here further points to an 2006). In the Biala Forest, the scuttle fly assem- overall tendency for both the dominants (species blages of pine plantations were characterised by ofrelative abundance 2 1%) and the characteristic higher values for the diversity and evenness indi- species on all the pine-plantation plots to be good ’ ces (H = 3.64 and J’ = 0.92) than those of old- colonizers, tolerant of abiotic stress, and mainly ’ growth stands (H = 3.47, J’ = 0.79). Further- generalists with multivoltine life cycle. Prescher more, the same three open-area species (M. (1992) and Buck (1997) observed the prevalence brevicostalis, M. verralli and Metopina Oligo— of the first dominant (Megaselia brevicostalis) neura) that show the highest fidelity to the pine also on each ofthe plots on open sites. It is charac- plantations ofthe Biala Forest, are also dominants teristic for scuttle fly assemblages to present two and characteristic species of pine plantations in peaks ofabundance (a lower one in the spring and the BialowieZa Primeval Forest and Tuchola Fo- a higher one in early autumn), the same holding rest (Durska 2006). true for plantations in the moist pine forest habitat Currently, little is known about the effective- of the BialowieZa Primeval Forest and Tuchola ness of collecting methods for phorid flies. Forest. The autumn rise and fall in abundances of Brown & Feener (1995) stated that Malaise traps Megaselz'a species is probably most connected are distinctly superior to pan traps in Southern with the abundance of fungi (Disney 1994, Hemisphere studies. My use here of the yellow Durska 1996, 200 1 a, 2006). Many scuttle fly spe- pan method reconfirrns the results of previous cies are known to be very sensitive to changes in work showing that Megaselz'a species make an their habitat, specially to climatic conditions extremely major contribution to assemblages of (Folgarait et al. 2007). During dry years, M. Phoridae (Disney 1994, Durska 1996, 2001a, brevicostalis reaches its abundance peak much 2006). Studies from Thunes et al. (2004), using a earlier than in years when humidity is typical motorised canopy fogger, also identified Mega— (Disney et al. 1981). selz'a as the best-represented genus, accounting In the present study I have shown that the as- for over 80% of individuals collected. During semblages of Phoridae present in Biala Forest four days of the experiment, only 91 individuals pine plantations are markedly different from of Phoridae were noticed. Among sixteen deter- those of its older pine stands. I attribute this dif- mined species, five species were present in the as- ference to the increasing shade cast as young trees semblages ofpine plantations in the Biala Forest. grow, and to the consequent replacement ofpho- In turn, individuals ofMegaselia spp. caught us- tophilous species ofopen areas by forest-dwellers ing Malaise traps in a wildfire site in dry pine fo- better adapted to the new conditions. ENTOMOL. FENNICA Vol. 20 ° Phoridae ofthe Biaia Forest, Poland 177

My earlier work documented dramatic de- Acknowledgements. My work on Phoridae has benefited from SYNTHESYS made available the Euro— clines in the abundance of the colonizers support by

early 7 pean Community Research Infrastructure Action under with shade increasing (Durska 1996, 2001a, the FP6 Structuring the European Area Programme AT- 2002, 2006). The availability of light can have a TAF 543 and SE-TAF 1833. I would like to thank James major impact on diversity. In general assem- Richards for English revision. For critical review of the manuscript and many valuable suggestions the author blages are first dominated by generalists of good wishes to thank two anonymous referees for their helpfiil that are for time’. Their dispersal ability ‘pressed comments. I am also grateful to Krzysztof Gagla, M. Sc., greater tolerance of abiotic stress is associated for his invaluable assistance with the segregation of the with search times that are more limited, ensuring material. that they “have to accept suboptimal choices” (Prinzing 2003; Fernandez & Costas 2004). References Taken together, the data I obtained during this Bankowska, R. & Garbarczyk, H. 1982: Charakterystyka and my previous studies suggest coherence terenéw badan oraz metod zbierania i opracowywania amongst Phoridae and the development processes materialow. In: Zoocenologiczne podstawy ksztalto- their assemblages pass through as moist pine fo- wania Srodowiska przyrodniczego osiedla mieszka- rests in the Polish Lowland undergo secondary niowego Bialoleka Dworska w Warszawie. Part I. Sklad i struktura terenu ektowa- succession 1996, 2001a, 2006, gatunkowy fauny proj

(Durska unpubl.). 7 nego osiedla mieszkaniowego. Fragmenta. Faunis- Morover, the same reaction the (characterised by tica 26 (1981): 17726. same group of dominant species and rather high Brown, B. V. & Feener, Jr. D. H. 1995: Efficiency of two species diversity) to disturbances (wildfire) ofthe mass sampling methods for sampling phorid flies in a open-area scuttle fly species has been obtained by (Diptera: Phoridae) tropical biodiversity survey. 7 Contributions in Science 459: 178. Prescher et al. (2002). A parallel response to dif- Buck, M. 1997: Untersuchungen zur okologischen Einni— ferent disturbances and (clear-cutting, grazing schung saprophager Dipteren unter besonderer burning) has also been found for spider and cara- Beriicksichtigung der Phoridae und Sphaeroceridae bid fauna (Gibson et al. 1992; Zulka et al. 1997; (Brachycera/Cyclorrhapha). Ph.D. thesis, Ulm Uni- Moretti et al. 2002; Fernandez & Costas 2004; versity, Germany. R. G. & E. M. 2005: Interaction bet- Huber & Baumgarten 2005). The high species di- Death, Zimmermann, ween disturbance and primary productivity in deter- on each that has versity present plot gone through mining stream invertebrate diversity. 7 Oikos 111: forest clearance and/or a wildfire reflects the at- 3927402. tendant habitat heterogeneity, and suggests a re- Disney, R. H. L. 1991: Family Phoridae. 7 In: Soos, A. & L. of Palaearctic Vo- silient response of scuttle fly assemblages to en- Papp, (eds), Catalogue Diptera. lume 7 (Dolichopodidae7Platypezidae): 1437204. vironmental stress. Resilience may be an indica- Akademiai Kiado, Budapeszt. tor of Manson et al. the stability, (2007) taking Disney, R. H. L. 1994: Scuttle flies: The Phoridae. Chap— term to mean of of “speed recovery community man & Hall, London. 467 pp. composition after perturbation”. The scuttle fly Disney, R. H. L. 2004: Species preferences for white ver— sus water for scuttle flies Phori— assemblage of a small patch opened up by distur- yellow traps (Dipt, 7 140: 317 bance differs in its composition in direct associa- dae). Entomologist’s Monthly Magazine 35. tion with how often such disturbances take place, Disney, R. H. L., Coluson, J. C. & Butterfield, J. 1981: A in the manner discribed the so-called “inter- by survey of the scuttle flies (Diptera: Phoridae) of mediate disturbance hypothesis” (IDH). This hy- upland habitats in Northern England. 7 Naturalist, Hull 106: 53766. pothesis could thus be tested using this group of R. H. L. & Durska, E. 2008: Conservation evalua— flies et al. 2004; Death & Zimmer- Disney,

(Roxburgh 7 tion and the choice of faunal taxa to sample. Bio- mann it is to use 2005). Accordingly, possible diversity and Conservation 17: 4497451. Phoridae, as a group of very high ecological di- Durska, E. 1996: The species composition and structure of versity, in monitoring of changes in terrestrial scuttle fly communities (Diptera: Phoridae) in mature in at habitats following disturbances, or as other indi- tree stands pine forests different stages ofhabitat degradation. 7 Fragmenta Faunistica 39: 2677285. cators within different ecological levels of orga- Durska, E. 2001a: Secondary succession of scuttle fly nization 1996, 2001a, 2006, Prescher et (Durska (Diptera: Phoridae) communities in moist pine forest al. 2002). Further investigations ofthese possibil- in BialowieZa Forest. 7 Fragmenta Faunistica 44: 8 17 ities remain necessary. 130. 178 Durska ' ENTOMOL. FENNICA Vol. 20

Durska, E.2001b: Zadrowate (Diptera: Phoridae). 7 In: Moore, P. D. 2005: Roots ofstability. 7Nature 437: 95 97 Gutowski, J. M. & Jaroszewicz, B. (eds), Catalogue of 961. the fauna of BialowieZa Primeval Forest: 2867289. Moretti, M., Conedera, M., Duelli, P. & Edwards, P.J. IBL, Warszawa. 2002: The effects ofwildfire on ground—active spiders Durska, E. 2002: The phenology of dominant scuttle-fly in deciduous forests on the Swiss southern slope ofthe (Diptera: Phoridae) species in the BialowieZa Forest. Alps. 7 Journal of Applied Ecology 39: 3217336. 7 Entomologica Fennica 13: 1237127. Gstman, 0., Kneitel, J. M. & Chase, J. M. 2006: Disturban— Durska, E. 2003: The phenology of Triphleba Rondani ce alters habitat isolation’s effect on biodiversity in species (Diptera: Phoridae) in moist pine forests in the aquatic microcosms. 7 Oikos 114: 3607366. BialowieZa Forest. 7 Entomologica Fennica 14: 1777 Prescher, S. 1992: Gkologie und Biologie der Diptera, ins— 182. besondere der Brachycera, eines klarschlammgedung— Durska, E. 2006: Diversity ofthe scuttle fly (Diptera: Pho- ten Ackerbodens. Ph.D. thesis, Braunschweig Univer— ridae) communities in the plantations ofmoist pine fo- sity, Germany. rests ofthe BialowieZa Primeval Forest and the Tucho- Prescher, S., Moretti, M. & Duelli, P. 2002: Scuttle flies la Forest (Poland). 7 Biodiversity and Conservation (Diptera, Phoridae) in Castanea sativa forests in the 15: 3857393. southern Alps (Ticino, Switzerland), with thirteen Fernandez Fernandez, M. M. & Salgado Costas, J. M. species new to Switzerland. Mitteilungen der Schwei— 2004: Recolonization of burnt pine forest (Pinus pi- zerischen Entomologischen Gesellschaft. 7 Bulletin naster) by Carabidae (Coleoptera). 7 European Jour— de la Societe Entomologique Suisse 75: 2897298. nal of Soil Biology 40: 47753. Prinzing, A. 2003: Are generalists pressed for time? An in- P. J R. J. W. & L. E. 2007: The test ofthe the time-limited model. Folgarait, ., Patrock, Gilbert, terspecif1c dispersal influence of ambient conditions and space on the pho— 7 Ecology 84(7): 174471755. nological patterns of a Solenopsis phorid guild in an Roxburgh, S. H., Shea, K. & Wilson, J. B. 2004: The inter— arid environment. 7 Biological Control 42: 2627273. mediate disturbance hypothesis: patch dynamics and Gibson, C. W. D., Hambler, C. & Brown V. K. 1992: mechanisms ofspecies coexistence. 7Ecology 85(2): Changes in spider (Araneae) assemblages in relation 3597374. to succession and grazing management. 7 Journal of Sorensen, T. 1948: A method of establishing groups of Applied Ecology 29: 1327142. equal amplitude in plant sociology based on similarity Goos, M. 1975: Influence of aphicides used in sugar—beet ofspecies content and its application to analyses ofthe plantations on arthropods. IV. Studies on the flying in— vegetation on Danish commons. 7 Biologiske Skrif— sects captured in yellow receptacles. 7 Polskie Pismo ter 5: 1734. Bulletin de K. J J Entomologiczne entomologique Pologne, Thunes, H., Skartveit, ., Gjerde, I., Stary, ., Solhry, T., Wroclaw 45: 4237144. [In Polish, with English sum- Fjellberg, A., Kobro, S., Nakahara, S., zur Strassen, mary]. R., Vierbergen, G., Szadziewski, R., Hagan, D. V., H. S. 1966: Measurement of in J Horn, ,,overlap” comparati- Grogan Jr., W.L., Jonassen, T., Aakra, K., Anonby, ., ve ecological studies. 7 American Naturalist 100: Greve, L., Aukema, B., Heller, K., Michelsen, V., Ha— 4197124. enni, J.-P., Emeljanov, A. F., Douwes, P., Berggren, Huber, Ch. & Baumgarten, M. 2005: Early effects offorest K., Franzen, J., Disney, R. H.L., Prescher, S., Johan— regeneration with selective and small scale clear-cut— son, K. A., Mamaev, B., Podenas, S., Andersen, S., ting on ground beetles (Coleoptera, Carabidae) in a Gaimari, S.D., Nartshuk, E., Srli, G. E. E., Papp, L., Norway spruce stand in Southern Bavaria (Hoglwald). Midtgaard, F., Andersen, A., von Tschimhaus, M., 7 Biodiversity and Conservation 14: 198972007. Bachli, G., Olsen, K. M., Olsvik, H., Foldvari, M., Lunau, K., Hofmann, N. & Valentin, S. 2005: Response of Raastad, J. E., Hansen, L. O. & Djursvoll, P. 2004: The the Hoverfly species Eristalis tenax towards floral dot arthropod community of Scots pine (Pinus silvestris guides with colour transition from red to yellow (Dip- L.) canopies in Norway. 7Entomologica Fennica 15: tera: Syrphidae). 7 Entomologica Generalis 27(3/4): 65790. 249725 6. Tanner, E. V. J. & Bellingham, P. J. 2006: Less diverse fo— Magurran, A. E. 1988: Ecological Diversity and its Meas- rest is more resistant to hurricane disturbance: eviden— urement. Princeton University Press, New Jersey. ce from montane rain forests in Jamaica. 7 Ecology Manson, N. W. H., Wilson, J. B. & Steel, J. B. 2007: Are 94: 100371010. alternative stable states more likely in high stress envi- Tschamtke, T. & Brandl, R. 2004: Plant-insect interactions ronments? Logic and available evidence do not sup— in fragmented landscapes. 7 Annual Review ofEnto— port Didham et al. 2005. 7 Oikos 116: 3537357. mology 49: 4057430. Matuszkiewicz, J., Degorski, M. & Kozlowska, A. 1993: Zulka, K.P., Milasowszky, N. & Lethmayer, Ch. 1997: Description ofthe plant association structure and soils Spider biodiversity potential of an ungrazed and gra- ofpine forest stands situated in five regions ofPoland. zed inland salt meadow in the National Park ,,Neusied— In: Species composition and structure of pine forests ler See—Seewinkel” (Austria): implications for mana- fauna in Poland. Part I. 7 Fragmenta Faunistica 36: gement (Arachnida: Araneae). 7 Biodiversity and 1 373 6. Conservation 6: 75788.